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Manuel Elías Gutiérrez & Lucía Montes Ortiz Estado actual del conocimiento de la diversidad del zooplancton (invertebrados) de agua dulce de la Península de Yucatán, utilizando la taxonomía integrativa Recibido:09/07/17 · Aceptado: 03/08/17 Manuel Elías Gutiérrez* Lucía Montes Ortiz El Colegio de la Frontera Sur Resumen Tradicionalmente se ha considerado que solo tres grupos se encuentran presentes en el zooplancton de agua dulce: rotíferos, cladóceros y copépodos. Sin embargo, existen muchos grupos más que potencialmente son parte de esta comunidad. En este documento se incluyen en el zooplancton a todos los invertebrados capaces de nadar e introducirse en una trampa de luz, lo que ampliará el universo de grupos estudiados. Por otro lado, la taxonomía integrativa involucra el uso de caracteres morfológicos, moleculares, geográficos, etc., para el reconocimiento de las especies. En este contexto, se incluirá a los zoopláncteres estudiados con métodos tradicionales y códigos de barras de la vida recolectados en la Península de Yucatán. Hasta el momento se han estudiado 1 768 ejemplares de 69 localidades incluyendo rotíferos, moluscos, colémbolos, ácaros acuáticos, cladóceros, ostrácodos, copépodos, malacostracos y quironómidos, comprendidos en 52 familias, 70 géneros y 132 especies. El estado curatorial de los especímenes hasta el nivel de especie es incompleto, por lo que el número total se incrementa hasta 205 posibles especies representadas por los índices de códigos de barras (BIN). La labor ha sido compleja, pues hay especies que pudieran ser nuevas para la ciencia o ampliaciones de ámbito. Por otro lado, también se han detectado formas extremas de una misma especie, como ocurrió con Bosmina tubicen, dentro del área de estudio, pudiéndose confundir fácilmente con B. huaronensis. Sin embargo, los códigos de barras indican una sola especie. Del mismo modo, especies morfológicamente muy similares pueden ser diferentes, como sucede con algunos ostrácodos. En conclusión, a pesar del avance logrado, es necesario continuar este trabajo, pues la comunidad de zooplancton puede ser un fuerte indicador de alteraciones antropogénicas en lugares todavía considerados prístinos, como la Laguna de Bacalar y la gran cantidad de cenotes que actualmente son atracciones turísticas. PALABRAS CLAVE: COI, códigos de barras de la vida, México, variabilidad, BOLD. *Correos electrónicos: [email protected], [email protected] 31 TEORÍA Y PRAXIS · ISSN 1870 1582 · NÚM. 25 · MAYO-AGOSTO 2018 · pp. 31-48 Present day kwnoledge on diversity of fresh water zoo- plancton (invertebrates) of the Yucatan Peninsula, using integrated taxonomy Recieved: 07/09/17 · Accepted: 08/03/17 Manuel Elías Gutiérrez* Lucía Montes Ortiz El Colegio de la Frontera Sur Abstract Traditionally it has been considered that only three groups are present in freshwater zooplankton: rotifers, cladocerans and copepods. However, there are many more groups that are potentially part of this community. In this document, all invertebrates capable of swimming and entering in a trap of light are included as zooplankton, which will expand the universe of groups studied. On the other hand, the integrative taxonomy involves the use of different characters morphological, molecular, geographical, etcetera for the recognition of the species. In this context, we will include zooplankters studied using traditional methods and DNA barcodes collected in the Yucatan Peninsula. So far, 1 768 specimens of 69 localities have been studied, including rotifers, mollusks, collem- bola, aquatic mites, cladocerans, ostracods, copepods, malacostraca and chironomids included in 52 families, 70 genera and 132 species. The curatorial status of specimens up to the species level is incomplete, so the total number is increased to 205 possible species represented by barcode index numbers (BINs). The work has been complex, as there are species that might be new to science or geographical range expansions. On the other hand, extreme forms of the same species have also been detected within the study area, as Bosmina tubicen, being easily confused with B. huaronensis. However barcodes indicate a single species. In the same way, morphologically similar species may be different, as it happens with some ostracods. In conclusion, despite the progress made, it is necessary to continue this work, as the zooplankton community can be a strong indicator of anthropogenic alterations in places still considered pristine, such as Laguna de Bacalar and the large number of cenotes that are currently tourist attractions. KEY WORDS: COI, DNA barcodes, Mexico, variability, BOLD. *E-mails: [email protected], [email protected] 32 TEORÍA Y PRAXIS · ISSN 1870 1582 · NÚM. 25 · MAY-AUGUST 2018 · pp. 31-48 Elías Gutiérrez / Montes Ortiz TEORÍA Y PRAXIS NÚM. 25 (2018: 31-48) Introducción Tradicionalmente se ha considerado que el zooplancton de agua dulce se en- cuentra constituido por tres grupos principales: rotíferos, cladóceros y copépo- dos. Sin embargo, existen otros grupos de organismos que también integran esta comunidad y que han sido escasamente contemplados como tales. A pe- sar de esto, algunos autores los han incluido en sus análisis como parte del zooplancton (Mwebaza-Ndawula, 1994, p. 261; Frost, Fisher, Klug, Arnott y Montz, 2006, p. 355), y es que varios de ellos exhiben claras adaptaciones a la vida planctónica, como los ácaros acuáticos (Modlin y Gannon, 1973, p. 223), los primeros estadios de quironómidos (Davies, 1976a, p. 42) y otros insectos (Janz, Weltje, Ebke y Dawo, 2016, p. 273), los ostrácodos (Deevey, Deevey y Brenner, 1980, p. 673), algunos decápodos e isópodos, entre otros. En este caso consideraremos como zooplancton todos aquellos organismos capaces de nadar en la columna de agua, aunque lo hagan por periodos cortos. Por otro lado, algunos de los sistemas dulceacuícolas de la Península de Yu- catán son calificados como ambientes extremos, debido a la presencia de aguas hipercarbonatadas, o de ciertos iones, como ocurre con la Laguna de Bacalar o el Cenote Azul (Perry, Velázquez-Oliman y Marín, 2002, p. 198). Por esta razón, y porque se trata de sistemas oligotróficos (Cervantes-Martínez, Mezeta-Barrera y Gutiérrez-Aguirre, 2009, p. 177; Smirnov y Elías-Gutiérrez, 2011, p. 215), se pen- saba que su riqueza de especies era limitada. Recientemente se han efectuado capturas con otros instrumentos, como trampas de luz (Elías-Gutiérrez et al., 2018), donde la diversidad de grupos encontrados ha sido sorprendente. En este trabajo se incluirán algunas capturas realizadas con esta nueva metodología, pero enfocándose principalmente al conocimiento taxonómico que se tiene de estos grupos. Sin embargo, todavía se desconocen otros, como se verá más adelante. Esta metodología, en conjunto con marcadores moleculares estan- darizados que se han adaptado ampliamente para la identificación de las espe- cies, conocidos como códigos de barras de la vida (Hebert, Cywinska, Ball y DeWaard, 2003, p. 313), ha permitido obtener interesantes resultados por Elías- Gutiérrez, Martínez-Jerónimo, Ivanova y Valdez-Moreno (2008, p. 5), donde también se incluyen algunas especies comunes de la Península de Yucatán. En el caso de México, este tipo de análisis ha sido un ejemplo pionero en el estu- 33 dio de los cladóceros (Bekker, Karabanov, Galimov y Kotov, 2016, p. 2) y los Estado actual del conocimiento de la diversidad del zooplancton (invertebrados) de agua dulce de la Península de Yucatán, utilizando la taxonomía integrativa restantes grupos del zooplancton (Elías-Gutiérrez, Martínez-Jerónimo, et al., 2008, pp. 8-14). La mayor parte de la información referente a códigos de barras que se ha generado en este país se encuentra en línea en la base de datos BOLD (Barcode of Life Data System, www.boldsystems.org). Actualmente, México ocupa entre el primer y tercer lugar en el nivel global en cuanto a especies secuenciadas con códigos de barras de los tres grupos principales de zooplancton dulceacuícola (rotíferos, cladóceros y copépodos), así como las primeras especies descritas considerando la taxonomía integrativa (Dayrat, 2005, pp. 408-409), donde se hace uso de caracteres morfológicos, moleculares, biogeográficos, etcétera. Entre las especies mexicanas descritas de esta forma destacan los cladóceros Leberis chihuahuensis Elías-Gutiérrez & Valdez-Moreno, 2008 (Elías-Gutiérrez y Valdez-Moreno, 2008, pp. 63-74) y Scapholeberis duranguensis Quiroz-Vazquez & Elías-Gutiérrez, 2009 (Quiroz-Vazquez y Elías-Gutiérrez, 2009, pp. 50-64), ambos del norte del país. Entre los copépodos sobresalen Leptodiaptomus gar- ciai (Osorio-Tafall, 1942), endémica del lago Alchichica (Montiel-Martínez, Ciros-Pérez, Ortega-Mayagoitia y Elías-Gutiérrez, 2008, p. 1080), y Mastigodi- aptomus patzcuarensis (Kiefer, 1938), del lago de Pátzcuaro (Gutiérrez-Aguirre, Cervantes-Martínez y Elías-Gutiérrez, 2014, p. 1). Respecto a la Península de Yucatán, las investigaciones en rotíferos son bas- tante limitadas y se ha trabajado solo de manera tradicional (García-Morales y Elías-Gutiérrez, 2007, p. 569; Sarma y Elías-Gutiérrez, 1999, pp. 187-196). El único trabajo donde se integran caracteres moleculares es el de García-Morales y Elías-Gutiérrez (2013, pp. 1097-1107), que abarca desde Yucatán hasta el al- tiplano de México, y donde se encuentra evidencia de la posible presencia de gran cantidad de especies crípticas, anteriormente consideradas formas, varie- dades o subespecies. Por otro lado, de los cladóceros hasta el momento se han descrito varias especies solo teniendo en cuenta aspectos morfológicos, como Antalona
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